Juniperus communis var. kelleyi, a new variety from North America

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Phytologia (August 2013) 95(3) 215 Juniperus communis var. kelleyi, a new variety from North America Robert P. Adams Biology Department, Baylor University, Box 97388, Waco, TX 76798, USA email Robert_Adams@baylor.edu ABSTRACT Recent molecular analysis of Juniperus communis, world-wide (Adams and Schwarzbach, 2012), has shown that taxa referred to as J. c. var. saxatilis in the eastern hemisphere and in North America are not the same taxon, but are found in two distinct clades. The name is correctly applied to the taxon in the eastern hemisphere, but the taxon referred to as var. saxitilis in North America is given a new name: Juniperus communis var. kelleyi R. P. Adams var. nov. in honor of a former student, Walter A. Kelley. Leaf terpene data is presented for J. c. var. kelleyi and other J. communis varieties as well as J. jackii. The leaf oil of var. kelleyi is dominated by α-pinene (56.5%), δ-3-carene (11.5%) and β-pinene (5.4%) and is very similar to the oil of J. c. var. depressa. Published on-line www.phytologia.org Phytologia 95(3): 215-221 (August 1, 2013). KEY WORDS: Juniperus communis var. kelleyi var. nov., nomenclature, DNA, leaf terpenes. Recently, Adams and Schwarzbach (2012) have shown that the North American taxon referred to as Juniperus communis var. saxatilis Pall. is not in the clade with J. c. var. saxatilis from Europe and Central Asia, but is actually most closely related to J. c. var. depressa from North America. To reflect these relationships, a new variety of Juniperus communis is described: Juniperus communis var. kelleyi R. P. Adams, var. nov. Fig. 1 Type: USA, Idaho, Blaine Co., on shore of Little Redfish Lake, 44 09.588' N, 114 54.372' W, 1997m, Adams 10892 (HOLOTYPE: BAYLU). Shrubs, similar to J. communis var. depressa, but differing in having curved to slightly curved leaves, with cross section concave and stomatal band 1.5-2 x width of green leaf margins, leaf blades free, 30 to 80 to the stem, seed cones about as long as leaves, seed cones ovoid, seed cones purple-blue when mature. Other specimens studied: TOPOTYPES: Adams 10890, 10891, 10893, 10894 at BAYLU. Juniperus communis var. kelleyi is common in the northwestern United States and B C, Canada (Fig. 2). In British Columbia and Alaska, var. kelleyi and var. depressa appear to intergrade. The new variety is named in honor of my former student, Walter A. Kelley, Ph. D. 1976, Colorado State University. Walt passed away, unexpectedly with a heart attack in Costa Rica, Dec. 31, 2010, while on one of his many trips to the rainforest with his wife, Jan. Walt worked on isozymes of Juniperus. Photo (right) shows Walt collecting samples of J. saltillensis (Nuevo Leon, Mexico) on a trip with Tom A. Zanoni and RPA. Walt's keen interest in plants and sense of humor will be missed.

216 Phytologia (August 2013) 95(3) Figure 1. Holotype of Juniperus communis var. kelleyi, Adams 10892 Figure 2. Distribution of Juniperus communis var. kelleyi. Table 3. Comparison of the leaf morphology of J. communis var. kelleyi, J. c. var. saxatilis and J. jackii. J. c. var. kelleyi J. c. var. depressa J. jackii Stomatal band width vs. green leaf margin (GM) 1.5-2x GM 1-1.5x GM 2-4x GM Leaf cross-section concave very concave concave, Leaf shape curved straight curved, boat shaped Leaf blades free, 30 to 80 free, 45 to 20 mostly appressed to stem Mature seed cones vs. cones about as cones much shorter than cones as long as or leaf length long as leaves leaves longer than leaves Seed cone shape ovoid ovoid elongated oval (ellisoid) esp. in immature cones

Phytologia (August 2013) 95(3) 217 The phylogenetic position of J. communis var. kelleyi is shown in Figure 3, where it is in a clade with the other J. communis varieties from North America. The only other member of section Juniperus in North America is J. jackii that is in a clade with J. mairei from China (Fig. 3). Figure 3. Bayesian tree for all taxa of Juniperus sect. Juniperus taxa. Numbers at the branch points are posterior probabilities (as percent). Adapted from Adams and Schwarzbach (2012). A minimum spanning network of the taxa of section Juniperus shows that J. communis var. kelleyi differs by only 2 MEs (mutational events) from var. depressa (Fig. 4), but var. kelleyi is a number of MEs different from the J. communis complex in Europe and central Asia (Fig. 4). A comparison of the leaf essential oil of var. kelleyi with var. saxatilis (Europe, Table 1) shows that these taxa differ in numerous components: α-pinene, sabinene, β-pinene, δ-3-carene, limonene, β- phellandrene, γ-terpinene, cis-sabinene hydrate, trans-sabinene hydrate, trans-thujone, terpinen-4-ol, myrtenol, citronellol, bornyl acetate, citronellyl acetate, neryl acetate, geranyl acetate, α-bisabolol, shyobunol, and 4 diterpenes. In fact, var. saxatilis (Europe, Table 1) seems to share a greater similarity in its oils with J. jackii, than with var. kelleyi (Table 1). The leaf oil of var. kelleyi is dominated by α- pinene (56.5%), δ-3-carene (11.5%) and β-pinene (5.4%) and is very similar to the oil of var. depressa (a similarity also seen in DNA sequence data, Figs. 3, 4).

218 Phytologia (August 2013) 95(3) Figure 4. Minimum spanning network (MSN) of the blue seed cone junipers (see notes Fig. 4). J. cedrus is the nearest of the red seed cone species. Note that c. var. kelleyi, Idaho, North America is separated by only 2 MEs from c. depressa and is not in the group with c. saxatilis from Europe - Central Asia. Adapted from Adams and Schwarzbach (2012). LITERATURE CITED Adams, R. P., P. S. Beauchamp, V. Dev and R. M. Bathala. 2010. The leaf essential oils of Juniperus communis L. varieties in North America and the NMR and MS data for isoabienol. J. Ess. Oil Res. 22: 23-28. Adams, R. P. 2000. Systematics of Juniperus section Juniperus based on leaf essential oils and RAPD DNA fingerprinting. Biochem. Syst. Ecol. 28: 515-528. Adams, R. P. and A. E. Schwarzbach. 2012. Taxonomy of Juniperus section Juniperus: Sequence analysis of nrdna and five cpdna regions. Phytologia 94: 280-297.

Phytologia (August 2013) 95(3) 219 Table 1. Comparison of the leaf oils of J. communis var. kelleyi with other varieties from North America and Europe and J. jackii. Taxa: J. jackii from serpentine, northwest California (jackii 1 ) and lava, Mt Hood, n Oregon (jackii 2 ); var. charlottensis, Queen Charlotte Island, BC (char); var. depressa, Guadalupita, NM (dep); var. megistocarpa, Magdalen Islands, Quebec (meg); var. kelleyi (Little Redfish Lake, ID); var. saxatilis, Switzerland, Europe (sax Eu); and var. communis, Stockholm, Sweden (com Sw). Data from Adams et al. (2010) and Adams (2000). Compounds in bold face appear to separate taxa. North America Europe KI Compound jackii 1 jackii 2 char meg dep kelleyi sax Eu com Sw 854 (E)-2-hexenal 0.2 1.0 0.4 0.1 0.1 0.3 1.2 0.7 926 tricyclene t t 0.1 0.1 0.1 0.1 t 0.3 931 -thujene t t t t t t 4.1 0.1 939 -pinene 16.1 18.9 59.3 58.5 53.9 56.5 14.1 56.8 953 -fenchene 0.3 0.6 0.1 t 1.0 0.5 0.1 0.3 953 camphene 0.3 0.6 0.6 0.6 1.0 0.5 0.2 0.6 954 thuja-2,4-diene - - 0.1-0.1 - - - 967 verbenene 0.3 0.3 - - - - - - 976 sabinene 0.1 0.3 0.3 0.3 0.3 0.3 32.8 0.7 978 1-octen-3-ol 0.1 - - t t - - - 980 -pinene 1.9 1.9 5.9 5.0 5.5 5.4 1.9 4.4 991 myrcene 3.2 3.2 4.8 3.9 4.1 4.5 5.0 5.2 997 ethyl hexanoate - - 0.1 - - - - 1001-2-carene 0.2 0.2 0.1 0.2 0.2 0.1 0.4 0.2 1005 -phellandrene 2.2 2.5 0.1 0.1 0.2 0.1 0.5 2.1 1011-3-carene 17.9 28.4 3.6 0.7 9.3 11.5 0.5 4.7 1018 -terpinene - - - 0.1 t - 1.9 t 1026 p-cymene 1.1 0.8 0.2 0.1 0.2 t 0.3 0.3 1031 limonene 6.6 0.5 1.9 20.4 2.6 2.1 6.7 5.1 1031 -phellandrene 13.4 9.2 2.9 1.0 2.5 3.1 0.6 8.9 1050 (E)- -ocimene 0.3 t - t t - 0.1 0.1 1057 amyl isobutyrate - - - - - - - 0.2 1062 -terpinene 0.1 0.1 0.1 0.1 0.1 0.1 3.4 t 1068 cis-sabinene hydrate - - - t - t 1.8 t 1088 terpinolene 3.2 4.4 1.0 0.5 1.4 1.8 3.0 1.1 1095 linalool - - 0.1 0.4 0.3 0.2-0.1 1097 trans-sabinene hydrate - - - - - - 1.3-1100 n-nonanal - - 0.1 - - - - - 1103 isoamyl-isovalerate - - - - - - t 0.1 1112 3-methyl-3-butenylisovalerate - - t t 0.1 0.1 - t 1114 trans-thujone (= β-thujone) - - - - - - 0.6-1121 cis-p-menth-2-en-1-ol 0.2 0.1 - t - t - t 1125 -campholenal 0.2 0.2 0.4 t 0.5 0.1 - t 1132 cis-limonene oxide 0.1 0.4-0.2 0.2 - - - 1133 cis-p-mentha-2,8-dien-1-ol - - - - - 0.1 - - 1139 trans-pinocarveol 0.2 0.3 0.4 t 0.5 0.1 - - 1141 camphor - - 0.2 - - 0.2 - - 1143 trans-verbenol 0.2 0.3 0.2 t 0.7 - - - 1147 3-methyl-2-butenylisovalerate - - - - 0.1 0.2 - t 1148 citronellal - - - 0.2 0.2 0.1 - - 1158 pinocarvone - - 0.1 - - - - - 1159 p-mentha-1,5-dien-8-ol 0.4 0.3 0.3-0.5 0.1 - t 1165 borneol - - 0.3 0.2-0.1 t 0.2 1172 cis-pinocamphone - - 0.1 - - 0.1 - - 1177 terpinen-4-ol 0.7 0.3 0.5 0.2 0.5 0.3 7.3 0.2 1176 m-cymen-8-ol - - 0.1-0.2 - - - 1179 naphthalene - - 0.1 0.2 - - 0.3 t

220 Phytologia (August 2013) 95(3) KI Compound jackii 1 jackii 2 char meg dep kelleyi sax Eu com Sw 1183 p-cymen-8-ol 0.3 0.3 0.1-0.2 - t t 1189 -terpineol 0.3 0.3 1.0 1.5 0.6 0.5 0.4 0.2 1190 methyl salicylate - - 0.1 - - 0.1 - - 1194 myrtenol 0.4 0.3 0.3 0.5 0.5 0.3 - - 1204 verbenone 0.3 0.5 0.3-0.3 t - t 1217 trans-carveol 0.4 t 0.3 t 0.2 t - - 1223 citronellol - - 0.1 0.3 0.5 0.2 - - 1235 methyl thymol 0.2 0.2 - t - 0.1 0.1-1239 carvone - - 0.1 - - 0.1 - - 1249 piperitone - - - t 0.4 0.2 - t 1257 methyl citronellate - - 0.2 t 0.1 0.3 - t 1285 bornyl acetate 0.5 0.5 1.0 0.5 0.6 0.7 0.2 0.9 1291 trans-verbenyl acetate - - - t - - - - 1292 (E,Z)-2,4-decadienal - - t t - t - - 1293 methyl myrtenate 0.2 0.5 - - - - - - 1302 -terpinyl formate 1.0 1.5 1.0-0.2 0.3 - - 1312 citronellic acid - - - t t t - - 1324 myrtenyl acetate 1.6 2.7 1.2 1.1 1.1 1.0 - t 1332 cis-piperitol acetate - - - - - 0.1 - - 1365 cis-carvyl acetate - - 0.1 - - t - - 1350 -terpinyl acetate 0.9 5.8 t 0.2 1.7 0.6 0.5-1350 citronellyl acetate - - 0.1 t 0.3 0.1 - t 1359 neryl acetate - - 0.1 t 0.1 0.1 - - 1379 geranyl acetate - - 0.1 0.1 1.3 0.3 - - 1381 trans-myrtanyl acetate t t - - - - t - 1391 -elemene 0.3 0.1 0.2 t 0.1 0.1 t 0.2 1418 (E)-caryophyllene 0.4 t t - - 0.1 t 0.7 1448 cis-muurola-3,5-diene - - 0.1 - - - - - 1454 -humulene 0.5 0.2 0.1 - t 0.1 t 0.5 1465 cis-muurola-4(14),5-diene t t 0.1 - - - - - 1475 trans-cadina-1(6),4-diene - - 0.1 - - 0.1 - - 1477 -muurolene t t 0.1 t 0.1 0.1 t t 1480 germacrene D 4.1 1.1 0.3 0.1 0.2 0.6 0.4 0.7 1493 trans-muurola-4(14),5-dien - - 0.1 - - 0.1 - - 1493 epi-cubebol 0.3 t 0.2 - - - - t 1499 -muurolene 0.6 0.2 0.3 0.1 0.1 0.2 0.2 0.2 1503 germacrene A t t 0.1 t 0.1 0.1 0.2 0.1 1505 β-bisabolene - - 0.1 - - - - - 1513 -cadinene 1.2 0.4 0.4 0.1 0.2 0.3 0.4 0.2 1524 -cadinene 2.2 0.7 1.4 0.4 0.5 0.7 0.8 0.5 1538 -cadinene 0.2 0.1 0.1 t 0.1 0.1 t t 1549 elemol t - - - t t - t 1556 germacrene B 0.5 0.3 0.3-0.3 1.2 0.3 0.3 1561 (E)-nerolidol - - t 0.1 t t - - 1574 germacrene D-4-ol 0.9 0.8 0.8 0.5 1.0 1.2 1.8 0.8 1577 spathulenol - - 0.1 0.1-0.1 - t 1607 -oplopenone - - 0.1 t - 0.1 - - 1581 caryophyllene oxide 0.2 t - - - - - t 1594 salvial-4(14)-en-1-one 0.1 - - - - - - - 1606 humulene epoxide II t t - - 0.1 - - t 1627 1-epi-cubenol 1.5 t t - - t t t 1640 epi- -cadinol 0.7 0.3 0.3 0.1 0.2 0.2 0.5 t 1640 epi- -muurolol 0.8 0.3 0.3 0.1 0.2 0.3 0.5 0.4 1645 -muurolol 0.4 0.1 0.1 t 0.1 0.1 0.1 t 1653 -cadinol 2.0 1.1 0.9 0.3 0.7 0.8 1.3 0.5 1685 -bisabolol - - 1.0 t 0.7 0.2 - - 1685 germacra-4(15),5,10(14)- trien-1-al 0.3 - - - - - - t 1688 shyobunol t t 0.2-0.1 0.3-0.7

Phytologia (August 2013) 95(3) 221 KI Compound jackii 1 jackii 2 char meg dep kelleyi sax Eu com Sw 1714 (2E,6Z)-farnesal - - - t - - - - 1722 (2Z,6E)-farnesal - - - 0.3 - - - - 1742 (2E,6E)-farnesal - - - t - - - - 1806 nootkatone - - 0.1 - - - - - 1933 cyclohexadecanolide 0.1-0.1 - - - - - 1968 sandaracopimara-8(14), 15-diene - t - - - - - - 2022 abieta-8,12-diene - t - - - - - - 1989 manoyl oxide 0.2 0.3 - - - - 0.1-2055 abietatriene 0.3 0.6 0.1 - - - 0.2-2056 manool 0.6 0.7 - - - - - - 2080 abietadiene - 1.1 - - - - 0.4-2106 isoabienol 0.2 0.9 - - - - 0.1-2331 trans-ferruginol t 0.2 - - - - - - KI = Kovat's Index on DB-5(= SE54) column. *Tentatively identified. Compositional values less than 0.1% are denoted as traces (t). Unidentified components less than 0.5% are not reported.